iforest ifor0210-0010001 battisti

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Review Article i Forest Biogeosciences and Forestry

Climate change and insectdistribution

The effects of global warming on living or-ganisms have now been recognized from the

level of individual species to communities,most notably in the form of temperaturere-lated range shifts (Walther et al. 22, !ootet al. 2", #ic$ling et al. 2%&. 's thenumber of insects per unit area is inverselyrelated to latitude and elevation (peight etal. )***&, we may assume that the increaseof temperature would allow the spreading ofinsect species northward and upward, espe-cially for those species that have wideranges, as many forest pests have. This as-sumption is supported by fossil data relatedto the forest insect response to climaticchanges of the past. #igher damage and in-sect diversity was recorded during the global

warming which occurred during the +aleo-cene ocene transition, relative to otherperiods (Wilf abandeira )***&.

With mean global temperatures increasingover the past ) years by about ./01 and

proected to continue (olomon et al. 23&,widespread climaterelated changes in thebiosphere can be e4pected. There are variousways by which the insects may react to cli-

mate change (Williams iebhold )**5,'yres ombardero 2, #arrington et al.2), 6ale et al. 22, +armesan 2%&, andit seems reasonable to assume that an in-crease of temperature within the vital limitsof a species implies a faster development.The species ready to e4pand are those char-acterized by high growth potential, mul-tivoltinism and absence of diapause, whereasthose that could be restricted show slow de-velopment rate and long cycles. The reduc-tion of the period of time spent as a larva orpupa may improve survival, as these are thestages more subected to predation and othermortality factors (6ernays )**3&. The in-

crease in population density may in turn pro-mote a further e4pansion of the range. omespecies would be simply limited in their sur-vival at the southern edge of their range andwould shift the range northward. witchingto new hosts may occur among nonspecial-ist herbivores, and can be the first con-se7uence of the strong selection on colon-izers (#arrington et al. 2), tastny et al.2%&.

+armesan 8ohe (2"& have provided a7uantitative assessment of the biological im-pact of climatic change, using data from dif-ferent types of organisms, including insects.

This analysis concerned the spatial (rangeshift& and phenological (advancement of

spring events& data, averaging %.) $m9decadeand 2." days9decade, respectively. ightypercent of the studied species (n : ;";&showed a consistent range shift and /3< an

advancement of spring events, such asflowering or migration.

#owever, the response of insects to climat-ic change may not always be linear (6ale etal. 22, =aston 2"&. >or e4ample, the de-velopmental stages of the insects can be dif-ferentially affected by the climate change,i.e. the growth can be accelerated by highertemperature, but at the same time the lengthof diapause may be e4tended.

Those insects developing without winterdiapause, which are active during this seasonand are protected from the low temperature,are the best candidates for range e4pansion ifthe winter temperature maintain the current

increasing trend (inclair et al. 2"&. Theincrease in winter temperature is a $ey factorfor the survival of the lepidopteranAtalopedes campestris in the new coloniza-tion areas (1rozier 2;&. ' good e4ampleconcerning a forest pest is the case of thepine processionary moth Thaumetopoeapityocampa(6o4 ),>ig. )&, reported also inthe last ?+11 report (!osenzweig et al.23&.

#owever, most forest insects of temperateregions have a winter diapause, which insome cases can last several years. Temperat-ure plays a maor role in the induction and

maintenance of this diapause. 'n increase ofthe temperature would modify the inductionand maintenance of the diapause, involvingchanges, which could affect the developmentof the insect, ma$ing predictions about popu-lation dynamics 7uite unreliable. Two e4-amples are reported here, which illustratehow high temperature during the larval de-velopment has caused lower diapause rateand higher damage by the spruce webspin-ning sawfly Cephalcia arvensis(6o4 2&, andhow high winter temperature has disruptedthe maintenance and termination of the eggdiapause in the larch bud moth Zeirapheradiniana(6o4 "&, causing a poor synchroniza-

tion with the host and the absence of an e4-pected outbrea$.

' different situation is presented by speciesthat are already adapted to the cold environ-ments, such asAglais urticae.These wouldprobably undergo a restriction of the range ifthey become limited at their southern bound-ary by increasing temperature (6ryant et al.)**3&.

>or insects that are heavily dependent on afavourable synchronization between budbrea$ing and hatching, such as the wintermoth Operophtera brumata, it appears thatthere may be a compensation between a

faster egg spring development and a delayedpupation in autumn, both triggered by an in-

SISEF http://www.sisef.it/iforest/ 1 iForest (2008) 1: 1-

* Dipartimento Agronomia Ambientale eProduzioni Vegetali, Universit di Padova,Via Universit 16, Agripolis, 3500 !egnaro"PD#, $tal%

&mail' Andrea (attisti"andrea)battistiunipd)it#)

!it"tio#' (attisti A, 00+) orests and

-limate -.ange / lessons rom inse-ts)iorest 1' 1/5 online' eb +, 00+2 U!'.ttp'44)sise)it4iorest4

Forests "#$ cli%"te ch"#&e - lesso#s fro%

i#sects

'"ttisti A

Astr"ct: *he cli%"te ch"#&e %"+ i#$irectl+ "ffects the forest ecos+ste%sthro,&h the "ctivit+ of ph+toph"&o,s i#sects. *he cli%"te ch"#&e h"s ee#cl"i%e$ to e respo#sile of the r"#&e ep"#sio# #orthw"r$ "#$ ,pw"r$ ofsever"l i#sect species of #orther# te%per"te forests "s well "s of ch"#&es i#the se"so#"l phe#olo&+. Sever"l p"pers h"ve $e"lt with the pre$ictio# of the%ost liel+ co#se,e#ces of the cli%"te ch"#&e o# the ph+toph"&o,s i#sectsi#cl,$i#& so%e of the %ost i%port"#t forest pests. I#cre"se$ levels of !2 i#the "t%osphere i#volve "# i#cre"se of the !/ "l"#ce of the pl"#t tiss,eswhich i# t,r# res,lts i# " lower foo$ ,"lit+ for %"#+ $efoli"ti#& i#sects. So%ei#sects respo#$ + i#cre"si#& the level of le"f co#s,%ptio# "#$ co#se,e#tl+the $"%"&e to the tree where"s others show hi&her %ort"lit+ "#$ lower per3for%"#ce. *he level of pl"#t che%ic"l $efe#ces %"+ "lso e "ffecte$ + "ch"#&e of !2. *he te%per"t,re is "ffecti#& either the s,rviv"l of the i#sects

which "re "ctive $,ri#& the col$ perio$ s,ch "s the pi#e processio#"r+ %othor the s+#chro#i4"tio# %ech"#is% etwee# the host "#$ the herivores "s i#the c"se of the l"rch ,$ %oth. A# i#cre"se of te%per"t,re %"+ "lter the%ech"#is% + which the i#sects "$5,st their c+cles to the loc"l cli%"te ($i"3p",se) res,lti#& i# f"ster $evelop%e#t "#$ hi&her fee$i#& r"te "s i# the c"seof the spr,ce we-spi##i#& s"wfl+ o,tre"s i# the So,ther# Alps.

6e+wor$s: te%per"t,re phe#olo&+ r"#&e host-i#sect rel"tio#ships


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Battisti A - iForest 1: 1-

crease of temperature (6use =ood, )**%&.Therefore, phenology is not affected, allow-ing 6ale et al. (22& to conclude that the ef-fects of the global warming would not be soevident in some species.

>inally, the natural enemies of forest in-sects may be affected by the temperature

change in different directions or e4tent. Thee4pansion of the host may not be promptlyfollowed by that of its enemies, as in thecase of the pine processionary moth (@ovi etal. in press&, or the synchronization between

host and parasitoid may not be maintainedunder new temperature conditions. Thisseems to be the case of a parasitoid of thewinter moth Operophtera brumata, which iseffective at low elevation but that is almostabsent at high elevation (Aersla$e et al.)**%&.

'll the e4amples cited above illustrate howinsects may react to the climate change,however they also have a great potential todevelop physiological and behavioural ad-aptations, which may improve their fitness

under new conditions. This would ultimatelylead to the formation of genetically differen-tiated populations and possibly new species,especially when the climatic change is asso-ciated with range e4pansion and host switch.

Climate change and host-insectinteractions

The capacity of an herbivore insect to com-plete its development depends on the adapta-tion to both, the environmental conditionsand the host plant. The plant may respond tochanges in temperature by varying its growth

iForest (2008) 1: 1- 2 SISEF http://www.sisef.it/iforest/

Fig. 1 gg batch of the pine proces-sionary moth Thaumetopoeapityocampaon a twig of!inus sylvestrisin the e4pansion outbrea$ area of Bont'vic ('osta&, at the unusual elevationfor an alpine population of );5 m. The

insect shows high performance and al-most no mortality due to the absence ofthe main natural enemiesC a genetic ana-lysis has shown that the population ori-ginated from different sources, suggest-ing that female moths are flying overbigger distances than previously be-lieved (alvato et al. 22&.

Box 1 - The pine processionarymoth Thaumetopoea

pityocampa

The winter pine processionary moth,Thaumetopoea pityocampaoffers a pos-sibility to test for the effects of globalwarming on an insect population over awide area of the Bediterranean basinand southern parts of urope, where it isthe most important pest of pine forests(!inus spp.&. ?ts geographic range lieswithin precise limits of elevation and lat-itude (DEmolin )*%*&, primarily as afunction of the average winter temperat-ures. 6ecause the larvae are oligophag-ous, potentially feeding on all !inusspp., but also on Cedrusspp. and the in-troduced !seudotsuga men"iesii, hostplant distribution does not restrict the

present range of the insectC many usualor potential host species grow in areaswhere the insect is absent. 1onse7uently,if the climatic conditions become fa-vourable in higher latitudes or at higherelevations, the insect may e4pand itsrange to these areas, often coupled withhost switching (6attisti et al. 25,!obinet et al. 23&. This relative im-portance of temperature over bioticfactors in defining the geographic distri-bution ma$es the moth a particularlysuitable model to study the range shift inrelation to global warming (6uffo et al.23&.

'n important forest pest in many areas,the moth has shown in the last decades asubstantial e4pansion of the outbrea$area both northward and upward(#uchon DEmolin )*3), #odar et al.2", 6attisti et al. 25&, aggravated bye4treme climatic events such as the sum-mer of 2" (6attisti et al. 2%&. Thishas resulted in high attac$ rates in areaspreviously largely unaffected by the in-sect (tastny et al. 2%&. The case de-serves special interest for the implica-tions it may have on the management ofuropean forests and plantations, as well

as on ornamental trees.

Fig. 2 alvation logging after repeated defoliation of the spruce webspinning sawflyCephalcia arvensisin Forway spruce stands of 1ansiglio +lateau (outhern 'lps&. This wasthe first outbrea$ of this insect ever recorded in the 'lps, where it is endemic.


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Forests and climate change - lessons #rom insects

rate, as it is commonly observed along latit-udinal or altitudinal gradients, when the con-ditions become progressively less favorableand the herbivores are more limited in the

hostGs e4ploitation. 'n e4ample is given bythe willow psyllid (Cacopsylla spp.&, whichis restricted to feed on a low number of spe-cies or types of plant tissues as the rangeedge is approached (#od$inson )**3, #ill etal. )**/&.

The change of temperature, which pro-motes the e4pansion of the insectGs range,may also involve a new association betweena herbivore and its host, as it has been shownby the pine processionary moth attac$ing themountain pine (!inus mugo& in the southern'lps. The large outbrea$s observed in thee4pansion areas on the new hosts may be e4-plained either by the high susceptibility of

the hosts or by the inability of natural en-emies to locate the moth larvae on an unusu-al hosts or environment (#odar et al. 2",tastny et al. 2%&.

'n elevated concentration of 1H2may af-fect the performance of phytophagous in-sects through the modification of the nutri-tional properties of the host plant (Iones etal. )**/, #unter 2), Anepp et al. 25&.'s 1H2is the main carbon source for photo-synthesis, its increase could alter thecarbon9nutrient balance of plants, increasingthe 19F ratio and thus diluting the nitrogencontent of the tissues. #owever, the responseof plants to increased 1H2 varies among spe-cies. ' high concentration causes an increase

of tannins in the leaves of birch, poplar andmaple, but not in the eastern white pine(!inus strobus& (!oth indroth )**;&.

The first reaction e4pected from herbivores

to the increase of the 19F ratio is compensat-ory feeding, in other words they should eatmore to accumulate enough nitrogen fortheir development. Thus, plant damage may

increase, but plant biomass could remainstable if we assume that the plants e4posedto high 1H2 grow more. +hytophagous in-sects may also develop adaptations to over-come higher 19F ratios, such as the pinesawfly$eodiprion lecontei, which shows an

increase in the efficiency of nitrogen utiliza-tion when reared on plants treated with high1H2 concentration (Williams et al. )**;&.#owever, other insect species seem unableto compensate the lower nutritional 7ualityof the plants by increasing the efficiency ofnutrient utilization (6roo$s Whitta$er)***, tiling et al. )***&. The e4perimentsof indroth et al. ()**"&, on three species ofsaturnid moths, show that the performance ofthe caterpillars is only marginally affectedwhen the nitrogen content of the leaves is re-duced by 2"< and the 19F ratio increasedby )"2/ig. 2&, during which the populations developedan annual life cycle and grew e4ponentially, causing repeated defoliations, which ulti-mately caused tree death over hundreds of hectares (6attisti et al. 2&. The most li$elyreasons for such a change in the life cycle of the insect have been e4plored through ananalysis of the local climate, which showed that the years preceding the outbrea$ werecharacterized by an abnormally warm and dry weather during the feeding period of thelarvae. We hypothesized that favourable conditions promoted the survival and speededup the development, ma$ing it possible to pupate when soil temperature was highenough to start pupation immediately, s$ipping in this way from the e4tended diapause.

ater, an e4periment showed that the soil threshold temperature for the induction of thee4tended diapause was about )2J1 (6attisti )**;&, well below the values recorded in theforest at the beginning of the outbrea$. The sudden increase of the population densitywas not 7uic$ly followed by that of natural enemies, which were unable to limit popula-tion growth (6attisti et al. 2&. ?t seems li$ely that the increase of the temperature inIune and Iuly of )*/"/5 is the maor factor promoting the outbrea$s, as they occurredsimultaneously at two sites at a distance of about ) $m, through switching the insectsto an annual generation.

Fig. 3 arch stand at timberline in the astern Dolomites, where the cyclic outbrea$s of thelarch bud mothZeiraphera dinianaare less regular and of variable intensity, fitting the mod-el of the easterly directional population waves created by 6ornstad et al. (22&, based onthe "5year dataset of 6altensweiler !ubli ()***&.


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!oth indroth ()**5& did not find higherparasitism by the hymenopteran Cotesiamelanoscelaon the larvae of%ymantria dis&par raised at high 1H2, whereas tiling et al.()***& found higher mortality of oa$ leafminers by parasitoids on two species of oa$

('uercus myrti#olia,'. geminata& grown athigh 1H2level.aboratory or greenhouse e4periments

provide valuable data, but it is difficult to de-

rive conclusions applicable to the natural en-vironment. >or e4ample, high 1H2levels are$nown to increase the temperature and, in-directly may affect the hostherbivore inter-action.

Dury et al. ()**/& showed that an increase

of "J1 of the temperature might lead to thesame effects of an increase of 1H2 (decreaseof nitrogen, increased of condensed tannins&on oa$ leaves. #owever, an increase of tem-perature may enhance the feeding of theherbivore and thus compensate for the negat-ive effects of a lower food 7uality. 'n e4per-iment that tested simultaneously the effectsof different levels of 1H2, nitrogen and tem-perature on the monoterpene production of!seudotsuga men"iesii (itva$ et al. 22&,indicated that the synthesis of these defencecompounds was more affected by individualtree variability than by the treatments.

The response of herbivore insects to in-

creased 1H2may also differ among the feed-ing guilds, as suggested by 6ezemer Iones()**/&. Defoliators are generally e4pected toincrease leaf consumption by about "'1 (>ree 'ir 1arbon nrichment& e4peri-ment carried out in Wisconsin (+ercy et al.22&, the activity of all guilds of herbivores,combined with the effect of increased ozone,may be compensated by the beneficial con-se7uences of enriched 1H2 on growth of

!opulus tremuloides.Bore research is clearly needed to ma$e re-liable predictions about the effects of climatechange on the relationships between theforest trees and phytophagous insects. =oodconceptual framewor$s, such as thecarbon9nutrient balance (6ryant et al. )*/"&and the growth9differentiation balance(#erms Battson )**2& theories are avail-able to interpret e4perimental results and toformulate new hypotheses. #opefully, thisunderstanding of the effects of climatechange on forest pests will enable us to ta$ethe necessary measures to counteract or mit-igate the possible negative conse7uences on

the forest ecosystems.

c!nowledgementsThis wor$ has been supported by the K 5 th

>+ LA51T22/52 proect M+ro-mothN (=lobal change and pine procession-ary mothO a new challenge for integrated pestmanagement& and by the proect M>orestBonitoringN of !egione del Peneto. ? than$Bichael tastny and doardo +etrucco Tof-folo for the useful discussions and for thehelp in screening the literature.

"eferences'yres B+, ombardero BI (2&. 'ssessing the

conse7uences of global change for forest disturb-

ance from herbivores and pathogens. cience of

the Total nvironment 2%2O 2%"2/%.

6ale I, Basters =I, #od$inson ?D, 'wmac$ 1,

6ezemer TB, 6rown PA, 6utterfield I, 1oulson

I1, >arrar I, =ood I=, #arrington !, #artley ,

Iones T#, indroth !, +ress B1, yrmnioudis

?, Watt 'D, Whitta$er I6 (22&. #erbivory inglobal climate change researchO direct effects of

rising temperature on insect herbivores. =lobal

1hange 6iology /O ))%.

6altensweiler W ()**"&. Why the larch budmoth

cycle collapsed in the subalpine larchcembran

pine forests in the year )** for the first time

since )/5. Hecologia *;O %2%%.

6altensweiler W, !ubli D ()***&. DispersalO an

important driving force of the cyclic population

dynamics of the larch bud moth, Zeiraphera

diniana =n. >orest now and andscape !e-

search 3;O ))5".

6attisti ' ()**;&. Poltinism and diapause in the

spruce webspinning sawfly Cephalcia arvensis.

ntomologia e4perimentalis et applicata 3O )5

))".

6attisti ', 6oato ', Basutti (2&. ?nfluence

of silvicultural practices and population genetics

on management of the spruce sawfly, Cephalcia

arvensis. >orest cology and Banagement )2/O

)5*)%%.

6attisti ', tastny B, Fetherer , !obinet1,

chopf ', !o7ues ', arsson (25&. 4pan-

sion of geographic range in the pine procession-

ary moth caused by increased winter temperat-

ures. cological 'pplications )5O 2/;2*%.

6attisti ', tastny B, 6uffo , arsson (2%&.

' rapid altitudinal range e4pansion in the pine

processionary moth produced by the 2" cli-matic anomaly. =lobal 1hange 6iology )2O %%2

%3).

6ernays ' ()**3&. >eeding by lepidopteran lar-

vae is dangerous. cological ntomology 22O

)2))2".

6ezemer TB, Iones T# ()**/&. +lantinsect herbi-

vore interactions in elevated atmospheric 1H2O

7uantitative analyses and guild effects. Hi$os /2O

2)2222.

6Qrnstad HF, +eltonen B, iebhold 'B, 6al-

tensweiler W (22&. Waves of larch budmoth

outbrea$s in the uropean 'lps. cience 2*/O

)2)2".

6roo$s =, Whitta$er I6 ()***&. !esponses of

three generations of a 4ylemfeeding insect,$eo&philaenus lineatus(#omoptera&, to elevated 1H2.

=lobal 1hange 6iology 5O "*5;).

6ryant I+, 1hapin > ???, Alein D! ()*/"&. 1ar-

bon9nutrient balance of boreal plants in relation

to vertebrate herbivory. Hi$os ;O "53"%/.

6uffo , 6attisti ', tastny B, arsson (23&.

Temperature as a predictor of survival of the pine

processionary moth in the ?talian 'lps. 'gricul-

tural and >orest ntomology *O %532.

6use ', =ood I= ()**%&. ynchronization of lar-

val emergence in winter moth (Operophtera bru&

mata .& and budburst in pedunculate oa$ ('uer&

cus robur .& under simulated climate change.

cological ntomology 2)O ""5";".

1rozier (2;&. Warmer winters drive butterfly

iForest (2008) 1: 1- SISEF http://www.sisef.it/iforest/

Box 3 - The larch bud mothZeiraphera diniana

The larch bud mothZeiraphera dinianais li$ely the best e4ample of a forest in-sect showing population cycles inurope, where it has been defoliatinglarge patches of larch stands in the 'lpsevery /) years for centuries (6altens-weiler !ubli )***&. This insect has anannual life cycle and spends the winteras an egg on the larch branches, startingto feed on the needles as soon as the budbrea$s. Bultiple factors seem to be in-volved in triggering the cycles, such asthe host plant 7uality, the natural en-emies, and the migration of the moths(6altensweiler !ubli )***, 6ornstadet al. 22, Turchin et al. 2"&.#owever, a disturbance probably relatedto climate change has caused the col-lapse of an outbrea$, which was startedin )*/* in the traditional occurrence area

in Kpper ngadine Palley (6altens-weiler )**"&. ?n the following years thelarval density decreased drastically and6altensweiler ()**"& was able to demon-strate that it was due to unusually highegg mortality, caused by repeated unfa-vourable weather conditions in winterand spring )*/**). ?n fact, it appearsthat weather is a critical factor for agood synchronization between bud burstand hatching of the larvae. ince theseconditions occurred in three successivegenerations, population growth was ef-fectively reduced and the cycle collapsedprematurely. ?n a recent analysis ofuropean temperature trend since )5(uterbacher et al. 2;&, it appears thatMthe winter )*/*9)** (RT:S2.;J1& andthe decade )*/* to )**/ (RT:S).2J1&were the warmest since )5. The period)*/* to )**/ was almost two standarderrors warmer than the second warmest(nonoverlapping& decade ()3"" to )3;2,RT:S.;5J1&, thus was very li$ely (*5orest ntomology "O )5")5*.

Anepp !=, #amilton I=, Bohan I, @angerl '!,6arenbaum B!, Deucia # (25&. levated

1H2 reduces leaf damage by insect herbivores in

a forest community. Few +hytologist )%3O 23

2)/.

Iones T#, Thompson I, awton I#, 6ezemer

TB, 6ardgett !D, 6lac$burn TB, 6ruce AD,

1annon +>, #all =, #artley , #owson =,

Iones 1=, Aampichler 1, Aandeler , !itchie

D' ()**/&. ?mpacts of rising atmospheric carbon

dio4ide on model terrestrial ecosystems. cience

2/O ;;);;".

Aersla$e I, Aruu$ 6, #artley , Woodin I

()**%&. Winter moth Operophtera brumata

(epidopteraO =eometridae& outbrea$s on cot-

tish heather moorlandsO effects of host plant and

parasitoids on larval survival and development.

6ulletin of ntomological !esearch /%O )55)%;.

indroth !, Ainney AA, +latz 1 ()**"&. !e-

sponses of deciduous trees to elevated atmo-

spheric 1H2O productivity, phytochemistry and

insect performance. cology 3;O 3%"333.

itva$ B, 1onstable IP#, Bonson !A (22&.upply and demand processes as controls over

needle monoterpene synthesis and concentration

in Douglas fir V!seudotsuga men"iesii (Birb.&

>ranco. Hecologia )"2O "/2"*).

uterbacher I, Dietrich D, Uopla$i , =rosean B,

Wanner # (2;&. uropean seasonal and annual

temperature variability, trends, and e4tremes

since )5. cience ""O );**)5".

Barchisio 1, 1escatti ', 6attisti ' ()**;&. 1li-

mate, soils and Cephalcia arvensisoutbrea$s on

!icea abiesin the ?talian 'lps. >orest cology

and Banagement %/O "35"/;.

+armesan 1 (2%&. cological and evolutionary

responses to recent climate change. 'nnual !e-

view of cology, volution and ystematic "3O

%"3%"*.

+armesan 1, 8ohe = (2"&. ' globally coherent

fingerprint of climate change impacts across nat-

ural systems. Fature ;2)O "3;2.

+ercy A, 'wmac$ 1, indroth !, Aubis$e

B, Aopper 6I, ?sebrands I=, +regitzer A,

#endrey =!, Dic$son !, @a$ D!, H$sanen7 ,

ober I, #arrington !, Aarnos$y D> (22&.

'ltered performance of forest pests under atmo-

spheres enriched by 1H2 and H". Fature ;2O

;";3.

!obinet 1, 6aier +, +ennerstorfer I, chopf ',

!o7ues ' (23&. Bodelling the effects of cli-

mate change on the potential feeding activity ofThaumetopoea pityocampa (Den. chiff.&

(ep., Fotodontidae& in >rance. =lobal cology

and 6iogeography VdoiO ).))))9.);%%/2"/.-

2%."2.4.

!oot T, +rice IT, #all A!, chneider #,

!osenzweig 1, +ounds I' (2"&. >ingerprints

of global warming on wild animals and plants.

Fature ;2)O 53%.

!osenzweig 1, 1asassa =, Aaroly DI, ?meson ',

iu 1, Benzel ', !awlins ', !oot T, eguin

6, Tryanows$i + (23&. 'ssessment of ob-

served changes and responses in natural and

managed systems. ?nO 1limate 1hange 23O ?m-

pacts, 'daptation and Pulnerability. 1ontribu-

tion of Wor$ing =roup ?? to the >ourth 'ssess-ment !eport of the ?ntergovernmental +anel on

1limate 1hange (+arry B, 1anziani H>,

+aluti$of I+, van der inden +I, #anson 1 eds&.

1ambridge Kniversity +ress, 1ambridge, KA,

pp. 3*)").

!oth A, indroth ! ()**;&. ffects of 1H2me-

diated changes in paper birch and white pine

chemistry on gypsy moth performance. Hecolo-

gia */O )"")"/.

!oth A, indroth ! ()**5&. levated atmo-

spheric 1H2 effects on phytochemistry, insect

performance and insect parasitoid interactions.

=lobal 1hange 6iology )O )3"/2.

alvato +, 6attisti ', 1oncato , Basutti , +atar-

nello T, @ane (22&. =enetic differentiation in

the winter pine processionary moth (Thaumeto&

poea pityocampa - (il)insonicomple4&, inferred

by '>+ and mitochondrial DF' mar$ers. Bo-

lecular cology ))O 2;"52;;;.

inclair 6I, Pernon +, Alo$ 1I, 1hown

(2"&. ?nsects at low temperaturesO an ecologic-

al perspective. Trends in cology and volution)/O 2532%2.

mith +#D, Iones T# ()**/&. ffects of elevated

1H2on the chrysanthemum leafminer, Chromat&

omyia syngenesiaeO a greenhouse study. =lobal

1hange 6iology ;O 2/32*).

olomon , Lin D, Banning B, 'lley !6,

6erntsen T, 6indoff F, 1hen @, 1hidthaisong

', =regory IB, #egerl =1, #eimann B, #ewit-

son 6, #os$ins 6I, Ioos >, Iouzel I, Aattsov P,

ohmann K, Batsuno T, Bolina B, Ficholls F,

Hverpec$ I, !aga !, !amaswamy P, !en I, !us-

ticucciB, omerville !, toc$er T>, Whetton +,

Wood, Wratt D (23&. Technical ummary. ?nO

1limate 1hange 23O The +hysical cience

6asis. 1ontribution of Wor$ing =roup ? to the

>ourth 'ssessment !eport of the ?ntergovern-

mental +anel on 1limate 1hange (olomon ,

Lin D, Banning B, 1hen @, Bar7uis B, 'veryt

A6, Tignor B, Biller # eds&. 1ambridge Kni-

versity +ress, 1ambridge, Knited Aingdom and

Few 8or$, F8, K'.

peight B!, #unter BD, Watt 'D ()***&. co-

logy of insectsO concepts and applications. 6lac$-

well cience, H4ford, KA.

tastny B, 6attisti ', +etrucco Toffolo ,

chlyter >, arsson (2%&. #ost plant use in

the range e4pansion of the pine processionary

moth, Thaumetopoea pityocampa. cological

ntomology")O ;/);*.tiling +, !ossi 'B, #ungate 6, Di$stra +,

#in$le 1!, Anott WB, Dra$e 6 ()***&. De-

creased leafminer abundance in elevated 1H2O

reduced leaf 7uality and increased parasitoid at-

tac$. cological 'pplications *O 2;2;;.

Turchin +, imon F, Wood +, Aendall 6, Bur-

doch WW, >ischlin ', 1asas I, Bccauley ,

6riggs 1I (2"&. Dynamical effects of plant

7uality and parasitism on population cycles of

larch budmoth. cology /;O )23)2);.

Walther =!, +ost , 1onvey +, Benzel ',

+armesan 1, 6eebee TI1, >romentin IB,

#oegh=uldberg H, 6airlein > (22&. cologic-

al responses to recent climate change. Fature

;)%O "/*"*5.Wilf +, abandeira 11 ()***&. !esponse of plant

insect associations to +aleoceneocene warm-

ing. cience 2/;O 2)5"2)5%.

Williams DW, iebhold 'B ()**5&. #erbivorous

insects and global change potential changes in

the spatialdistribution of forest defoliator out-

brea$s. Iournal of 6iogeography 22O %%5%3).

Williams !, incoln D, Thomas !6 ()**;&.

oblolly pine grown under elevated 1H2 affects

early instar pine sawfly performance. Hecologia

*/O %;3).

@ovi D, tastny B, 6attisti ', arsson (2/&.

cological costs on local adaptation of an insect

herbivore imposed by host plants and natural en-

emies. cology (in press&.

SISEF http://www.sisef.it/iforest/ iForest (2008) 1: 1-

iforest ifor0210-0010001 battisti

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